A Taxonomic Review of the Dark-Winged Trumpeter Psophia Viridis (Aves: Gruiformes: Psophiidae)

Volume 49(41):547‑555, 2009

A taxonomic review of the Dark-winged Trumpeter Psophia viridis (Aves: Gruiformes: Psophiidae)

Marina Oppenheimer1,2 Luís Fábio Silveira1,3

Abstract

The Dark-winged Trumpeter, Psophia viridis (Gruiformes, Psophiidae) is a Brazilian endemic species and includes three subspecies: Psophia viridis viridis Spix, 1825; Psophia v. dextralis Conover, 1934, and Psophia v. obscura Pelzeln, 1857, as well as P. v. interjecta Griscom & Greenway, 1937, whose validity has been questioned by several authors. These taxa are allopatric in distribution along the south of the Amazon River, although the precise limits of their distribution still remain unknown. This complex has never been taxonomically reviewed and this work aims to test the validity of its taxa based on the Phylogenetic Species Concept. Morphometrical characters and plumage colour patterns were analyzed, and the distribution of the taxa was also revised. In this study, 108 specimens from 41 localities were examined (all types included), with each reliable literature-based locality being included in order to delimit the geographical distribution of the complex. Morphometrical data did not point out significant differences between the taxa, also showing no sexual dimorphism among them. Meanwhile, plumage characters showed consistent and distinct patterns for each of the taxa, except for P. v. interjecta, whose features indicated by authors as diagnosable are the result of individual variation. No clinal variation or intergradation were observed, even at regions close to the rivers headwaters, where supposedly populations could be in contact. It is suggested that the currently accepted subspecies be elevated to the species level, such as: Psophia viridis Spix, 1825, distributed in the Madeira-Tapajós interfluvium, P. dextralis, found in the Tapajós- Tocantins interfluvium, and P. obscura, which occurs from the right bank of the Tocantins River to the west of the State of Maranhão.

Keywords: Psophiidae; Psophia; Psophia viridis; Taxonomy.

Introduction The Gray-winged Trumpeter (Psophia crepitans) occurs north of the Amazon river, from French Guiana The genus Psophia Linnaeus, 1758 includes to Peru and Ecuador; the White-winged Trumpeter three species, all restricted to Amazon basin forests. (P. leucoptera) is found north and south of the Ama-

1. Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11.461, 05422-970, São Paulo, SP, Brasil; Seção de Aves, Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42.494, 04218‑970, São Paulo, SP, Brasil. 2. E‑mail: [email protected] 3. E‑mail: [email protected] 548 Oppenheimer, M. & Silveira, L.F.: Taxonomy of Dark-winged Trumpeter zon, from eastern Peru to the Madeira river in the west, Sharpe (1894) considered Psophia obscura to be and Bolivia. The Dark-winged Trumpeter (P. viridis), a full species, while Peters (1934) made it subordinate is a Brazilian endemic and also occurs south of the to P. viridis, as it is treated today. Griscom & Green- Amazon river, but east of the Madeira river to the way (1937) observed that P. v. interjecta has features state of Maranhão (Sherman, 1996). of both P. v. obscura and P. v. viridis, while intermedi- Depending on the author, the Dark-winged ate geographically between P. v. obscura and P. v. dex- Trumpeter may include three or four taxa. The nomi- tralis. Haffer (1974) stated that P. v. interjecta is indis- nate form occurs in the region between the Madeira tinguishable from P. v. dextralis and suggested that it and Tapajós rivers, and P. v. dextralis occurs between is an intergradation of P. v. dextralis and P. v. obscura, the Tapajós and Xingu rivers. The putative and de- and only subtly different from them, being possibly bateable P. v. interjecta occurs from the east bank of found in simpatry with both. Although he did not the Xingu to the west margin of the Tocantins river, synonymize the two, Haffer’s treatment (1974) was being replaced by P. v. obscura from the east bank of followed by many (e.g. Sherman, 1996), yet Pinto the Tocantins to western Maranhão (Peters, 1934; (1978) considered P. v. interjecta to be valid while Blake, 1977; Pinto, 1978; Sherman, 1996; Sick, recognizing that its diagnosis from the others remains 1997). uncertain. Spix (1825:66) described Psophia viridis based Since the original descriptions, the taxa belong- on a specimen from “Villa Nuova” (= Parintins, Ama- ing to the Psophia viridis complex have not been ex- zonas), the only island locality for Psophia. He de- tensively reviewed. Here, we address this question by scribes the purple iridescence in the lower neck, the examining all of the taxa involved to examine and green back, and calls attention to dark green scapulars, compare their morphological variation and geograph- whereas the greater wing-coverts are rusty. Psophia ob- ic distribution. scura Pelzeln, 1857 was the next to be described, based on three specimens collected by J. Natterer. The type locality given by Pelzeln (1857) was simply “Brasilia”, Material and Methods later changed by him to “Pará” (Pelzeln, 1871). Nat- terer (in Pelzeln, 1857) states that he collected three We analyzed 108 adult specimens of the Pso- specimens on January 1835. At that time he was in phia viridis complex from 41 localities, comprising Belém or nearby (Vanzolini, 1993), which suggests all described taxa (Appendix 1). These specimens are that the type locality should be Belém, as also sug- housed in the collections of Museu de Zoologia da gested by Hellmayr & Conover (1942) and Pinto Universidade de São Paulo, São Paulo (MZUSP), Mu- (1978). Pelzeln (1857) pointed that P. obscura differs seu Paraense Emílio Goeldi, Belém (MPEG), and Mu- from P. viridis by its brownish-green mantle and dis- seu Nacional, Universidade Federal do Rio de Janeiro, creet purple iridescence at the base of the neck. Rio de Janeiro (MNRJ). Type specimens of Psophia Psophia viridis dextralis Conover, 1934, was v. viridis (Zoologische Staatssammmlung München, described based on an adult male collected by A.M. Munich, Germany – ZSM B11) and P. v. dextralis Olalla on 02 December 1932 at Tauari, Tapajós river, (Field Museum of Natural History, Chicago, USA – Pará. The diagnostic characters of P. v. dextralis are FMNH 410480) were analyzed by us, while the type the greenish olive mantle and that the iridescence of specimens of, P. v. interjecta (Museum of Compara- the wings is almost absent and, when not, is greenish. tive Zoology, Cambridge, USA – MCZ 173207) and Three years after P. v. dextralis was described, P. v. in- P. v. obscura (Naturhistorisches Museum Wien, Wien, terjecta was described from an adult male from Cam- Austria – NHMW 39401) were analyzed from pho- etá, left (west) bank of the Tocantins river (Griscom tographs. Measurements of bill, wing, tail and tarsus & Greenway, 1937). Psophia v. interjecta was consid- length followed Baldwin et al. (1931). ered as a combination of characters from P. v. obscura Colors were determined following the catalogs and the nominate form. The only specimen used to of Smithe (1975; hereafter S) and of Munsell (1994; describe this taxon was different from two topotypic hereafter M). All specimens were analyzed under natu- pairs of P. v. dextralis available to the authors, which ral light. Plumage characters used included some that led them to consider P. v. interjecta to be a valid taxon were used in the past to separate taxa, and included: (Griscom & Greenway, 1937). Diagnosis is based on a) color of the mantle in the upper (proximal to the the purple iridescence of the wing and on the brown neck), intermediate, and lower portion (distal from upper part of the mantle, thus separating it from the neck); b) iridescence in the neck (presence – ab- P. v. obscura. sence); and c) purple in the wing (presence – absence). Papéis Avulsos de Zoologia, 49(41), 2009 549

Colors of other body parts were not used here due to portion of the mantle varied between Parrot Green the inherent variation found in museum specimens. (S 260) and Olive Green, while the proximal portion When not included with the specimens, the col- was Dark Green (S 162A). Again, the exception was lecting locations were determined following Paynter specimen MNRJ 9645, which has Very Dark Brown Jr. & Traylor (1991) and Vanzolini (1992). All loca- (M 7.5YR/2.5/3). This appears to be an individual tions were mapped with ArcView (ArcView 3.3 for variation since another specimen collected 70 km of Windows, ESRI, 2002). Map preparation also includ- this specimen (MZUSP 76728) is typical to the re- ed data from Spix, 1825; Pelzeln, 1857; Brabourne mainder of those birds. All specimens from this re- & Chubb, 1912; Snethlage, 1914; Naumburg, 1930; gion have iridescent purple on their wings and neck Peters, 1934; Conover, 1934; Griscom & Greenway, (Table 1; Figures 1-3a). 1937; Pinto, 1938, 1978; Hellmayr & Conover, A second group (n = 73), collected between the 1942; Gyldenstolpe, 1945; Schauensee, 1970; Blake, Tapajós and the Tocantins rivers, is very homoge- 1977; Graves & Zusi, 1986; Stotz, 1986; Oren, 1990, neous in color. The distal portion of the mantle is Ol- 1991; Novaes & Lima, 1991. ive Green (S 46), with two exceptions (MPEG 51281 and 51284), with the former being Leaf Green (S 146) and the latter being Greenish Olive (S 49). Two oth- Results and Discussion er specimens in this same series (MPEG 51282 and 51283) have the distal portion of the mantle Olive We found no morphometric differences be- Green, in agreement with all the other specimens. tween the sexes and among the described taxa (data The intermediate and proximal mantle of all but two not shown, available under request). Color patterns specimens are Very Dark Brown (M 7.5YR/2.5/3 and resulted in three well defined groups. Birds from /2). However, there are two exceptions: specimens the region between the Madeira and Tapajós rivers from Santana do Araguaia, Pará (MPEG 48495 and (24 specimens) have, with one exception, the distal 48496), have the entire mantle Leaf Green and two part of the mantle Parrot Green (S 160 and 260). specimens from Carajás, Pará (MPEG 37204 and The exception was specimen MNRJ 9645, which 37205) have the intermediate part of the mantle Leaf has this area Olive Green (S 47). The intermediate Green and Olive Green, respectively. Wings and neck

Figure 1: Pattern of the mantle coloration. Specimens from Madeira-Tapajós interfluvium (a, MZUSP 62339); Tapajós-Xingu interfluvium (b, MPEG 22098), Xingu-Tocantins interfluvium (c, MPEG 37970), and to the east of Tocantins river and west of Maranhão (d, MZUSP 43899). 550 Oppenheimer, M. & Silveira, L.F.: Taxonomy of Dark-winged Trumpeter

Figure 2: Pattern of the wing coloration. Specimens from Madeira-Tapajós interfluvium (a, MZUSP 62339); Tapajós-Xingu interfluvium (b, MPEG 22098), Xingu-Tocantins interfluvium (c, MPEG 37970), and to the east of Tocantins river and west of Maranhão (d, MZUSP 43899). iridescence is almost or entirely absent in all speci- rot Green and the proximal portion Dark Green. The mens analyzed (Table 1; Figures 1-3b, c). neck and wings have a conspicuous purple iridescent. The third group (n = 11), found from eastern Conover (1934) compared a specimen of Psophia Tocantins to Maranhão, has the distal and interme- viridis collected on the east bank of the Tapajós river diate portion of mantle consistently Dark Green (Tauari) with three specimens from Madeira river (in- (S 162A and 262), while the proximal portion is Dark cluding the type) and two P. obscura from the Natural Brown (M 7.5YR/3/2). Wings and neck iridescence History Museum of Wien to describe P. v. dextralis. is almost or entirely absent on this group (Table 1; Additionally, he refers to a specimen from the Ca- Figures 1-3d). maraipi River (Pará) that has more in common with Psophia viridis, according to Spix (1825), has a P. v. obscura than with P. v. dextralis, suggesting that green mantle and the lower neck is iridescent purple, specimen is perhaps transitional between P. v. dex- both considered to be diagnostic features for this spe- tralis and P. v. obscura (see below). Two topotypes in cies. The colors found in the holotype and in a topotype the MZUSP have the distal portion of the mantle Ol- (MZUSP 10938) are in agreement with the original ive Green, in agreement with the holotype (FMNH description and are constant in the specimens between 410480). Also, purple iridescence on the wings and the Madeira and Tapajós rivers. Also, P. viridis Spix, neck is almost absent. These characters are consistent 1825 can be separated from other Psophia by having in specimens from the area between the Tapajós-To- the distal and intermediate portion of the mantle Par- cantins rivers. Psophia viridis dextralis Conover, 1934

Figure 3: Pattern of coloration of the base of the neck. Specimens from Madeira-Tapajós interfluvium (a, MZUSP 62339); Tapajós- Xingu interfluvium (b, MPEG 22098), Xingu-Tocantins interfluvium (c, MPEG 37970), and to the east of Tocantins river and west of Maranhão (d, MZUSP 43899). Papéis Avulsos de Zoologia, 49(41), 2009 551

Table 1: Plumage patterns found on dark-winged trumpeters

Interfluvium Character To the east of the Madeira-Tapajós Tapajós-Xingu Xingu-Tocantins Tocantins river Mantle Distal portion Parrot Green (S 160) Olive Green (S 46) Olive Green (S 46) Dark Green (S 162A or (S 2600) e 262) Intermediate portion Parrot Green (S 260) Very Dark Brown Very Dark Brown Dark Green (S 262) and Olive Green (S 47) (M 7.5YR/2.5/3) (M 7.5YR/2.5/3) Proximal portion Dark Green (S 162A) Very Dark Brown Very Dark Brown Dark Brown (M 7.5YR/2.5/2) (M 7.5YR/2.5/2) (M 7.5YR/3/2) Iridescence Wing Purple Almost or entirely Almost or entirely Almost or entirely absent absent absent Lower neck Purple Almost or entirely Almost or entirely Almost or entirely absent absent absent has as unambiguous diagnostic characters the lower characters of the nominate form could be found in portion of the mantle Olive Green, intermediate and P. v. interjecta. upper parts of the mantle Dark Brown, and irides- Our results show that specimens from the puta- cence on the wings and neck discreet or absent. tive range of P. v. interjecta do not possess characters Griscom & Greenway described P. v. interjecta which can diagnose them from P. v. dextralis. Thus, based in a single specimen, and stated that this tax- according to morphological characters, P. v. interjecta on presented a combination of the characters from cannot be considered a valid taxon, and therefore it P. v. obscura and the nominate form. The characters of is an individual variation of P. v. dextralis. Some au- P. v. obscura observed on P. v. interjecta would be the thors have already recognized this, but also suggested discreet iridescence both on the neck and the wings that these may be intermediate between P. v. dextralis and the mantle darker than observed in P. v. obscura. and P. v. obscura (Blake, 1977; Pinto, 1978; Sher- Griscom & Greenway (1937) do not determine which man, 1996), but our analysis rejects this hypothesis.

Figure 4: Distribution of dark-winged trumpeters in Brazil. Asterisk refers to the type-locality of each taxa. Points with a central dot refer to records obtained from literature. Circles refers to specimens analyzed of Psophia viridis, triangles to P. dextralis (including P. v. interjecta), and squares to Psophia obscura. Brazilian States are Amazonas (AM), Pará (PA), Rondônia (RO), Mato Grosso (MT), Tocantins (TO) and Maranhão (MA). 552 Oppenheimer, M. & Silveira, L.F.: Taxonomy of Dark-winged Trumpeter

Table 2: Diagnostic characters for Psophia viridis, Psophia dextralis and Psophia obscura.

Mantle Taxon Lower Neck Wing Distal portion Intermediate portion Proximal portion P. viridis Parrot Green Parrot Green Dark Green Purple iridescence present Purple iridescence present and conspicuous and conspicuous P. dextralis Olive Green Very Dark Brown Very Dark Brown Discreet or absent Discreet or absent P. obscura Dark Green Dark Green Dark Brown Very much reduced or Very much reduced or absent absent

Although the original description of P. v. interjecta Psophia viridis Spix, 1825 includes extensive blue and iridescent wing coverts (“extensive blue apical spots to the wing-coverts”), only Type-locality: Parintins (“Villa Nuova”), Amazonas. one of the specimens analyzed (MPEG 37205) had this color on the wing coverts and therefore should be Diagnosis: distal and intermediate portions of the considered to be an individual variation as observed in mantle Parrot Green (S 160; S 260), and upper por- the mantle of some specimens from this same area. tion Dark Green (S 162A); purple iridescence of the Pelzeln (1857) described Psophia obscura based wings and neck present and conspicuous. on three individuals without designating a holotype, designating an adult male as the lectotype some years Distribution: Madeira-Tapajós interfluvium, with the later (NHMW 39401, Pelzeln, 1871). Pelzeln (1857) northern limit at its type-locality, Parintins (Amazo- noted that P. obscura differs from P. viridis due to its nas; MZUSP 10938), and the southern limit at Águas smaller bill and tarsus, discreet purple iridescence at do Guaporé (Rondônia, MNRJ 9644; Figure 4). the base of the neck and a brownish-green mantle. Here, we show that P. v. obscura can be separated both from P. v. viridis and P. v. dextralis based on the Dark Psophia dextralis Conover, 1934 Green (S 162A and S 262) color of the distal and intermediate portion of the mantle, being Dark Brown Psophia viridis interjecta Griscom & Greenway, 1937 (M 7.5YR/3/2) in the upper region. The purple iri- syn. jun. descence of the wings and the neck, when present, is less noticeable than in P. v. viridis. Type-locality: Tauari, Pará. We found five distinct and non-overlapping plumage characters that unambiguously diagnose Diagnosis: distinguished from the other taxa by pre- each one of the three groups of specimens established senting the distal portion of the mantle Olive Green above (see Table 2), and we also noted a lack of clinal (S 46), and the intermediate and proximal portion variation and putative hybrids. Helbig et al. (2002) of the mantle Very Dark Brown (M 7.5YR/2.5/3 e state that a taxon is diagnosable if individuals of the M 7.5YR/2.5/2). The purple iridescence of the wings same age and sex are different from individuals of the and the neck is discreet or absent. same age and sex of all of the other taxa by at least one qualitative difference, or if one category of age Distribution: between the Tapajós and Tocantins riv- and sex is separated by a complete discontinuity in at ers, with the northern limit at Portel (Pará, MPEG least one continuous and variable character (quanti- 40708 and MPEG 40709) and the southern limit at tative character) of the same category of age and sex Sete de Setembro river (Mato Grosso, MPEG 14781; of other related taxa. Based on morphological char- Figure 4). acters, our findings fulfill this definition and lead us to consider that the former Psophia viridis complex comprises three distinct species, which differ in their Psophia obscura Pelzeln, 1857 mantle color and in the iridescence of the wings and base of the neck. Psophia viridis interjecta Griscom & Type-locality: Belém, Pará (fide Hellmayr & Conover, Greenway, 1937 shows no diagnostic character and 1942). should be considered as a junior synonym of P. dextralis Conover, 1934. The diversity of dark-winged Diagnosis: distinguished from the other species by trumpeters is represented by the following species: the distal and intermediate portion of the mantle Papéis Avulsos de Zoologia, 49(41), 2009 553

Dark Green (S 162A and 262), being Dark Brown P. obscura is the urgent conservation of the remain- (M 7.5YR/3/2) in the proximal portion. The purple ing forested areas of the east of Pará and Maranhão iridescence of the wings and neck is very much re- (Oppenheimer, 2008). The creation and effective duced or absent in most specimens. protection of conservation units or private reserves on the remaining preserved forests in the Belém Distribution: Occurs to the east of Tocantins river and center of endemism is essential not only for this west to Buriticupu (Maranhão, MPEG 37338; Oren taxon but for the other endemic and threatened 1990, 1991). The northern limit is Belém, the type species in this region. locality, and the southern limit is Jacundazinho (Pará, MPEG 36328; Figure 4). Resumo These species of Dark-winged Trumpeters are allopatric. The large rivers of the Amazon basin seems Os jacamins-de-costas-verdes, Psophia viridis to be efficient barriers and may often prevent contact (Gruiformes, Psophiidae) são endêmicos da Amazônia between related populations, as partially observed brasileira e contam, atualmente, com três subespécies with these trumpeters. While complete isolation by reconhecidas: Psophia viridis viridis Spix, 1825; rivers is rarely observed (Haffer 1974), the differ- Psophia v. dextralis Conover, 1934, e Psophia ences in plumage could disappear clinally towards the v. obscura Pelzeln, 1857, além de P. v. interjecta headwaters, and the river would be less effective as Griscom & Greenway, 1937, cuja validade tem a barrier for dispersal, allowing contact between the sido questionada por diversos autores. Estes táxons populations. Despite the sampling of Dark-winged distribuem-se alopatricamente ao sul do rio Amazonas, Trumpeters are far from ideal, most of the specimens embora os limites precisos de suas distribuições sejam were collected along the rivers and we examined ainda pouco conhecidos. Os táxons deste complexo nunca specimens collected at the headwaters of some main passaram por uma revisão taxonômica, e o presente rivers south of Amazon. We were able to check the trabalho teve como objetivos testar a validade dos constancy of the characters especially on these areas mesmos. Com base no Conceito Filogenético de Espécie, and we did not see any plumage variations associ- os caracteres morfométricos e de colorido da plumagem ated with latitude, being conservative even at regions foram analisados, além de refinar a distribuição where the rivers are narrower and supposedly the geográfica dos táxons do complexo. Foram examinados taxa could be in contact. We found no evidence for 108 espécimes procedentes de 41 localidades, incluindo clinal variation, and color patterns appeared consis- todos os tipos, e foram considerados também os registros tent along the species’ distribution. The Xingu river confiáveis de literatura para delimitar-se a distribuição did not isolate (at least from a morphological point geográfica. Os dados morfométricos não indicaram of view) populations of the Dark-winged Trumpet- diferenças significativas entre os táxons, não apontando ers while the Madeira, Tapajós and Tocantins rivers também qualquer dimorfismo sexual. Entretanto, a did, separating populations and eventually prevent- análise dos caracteres de plumagem evidenciou padrões ing gene flow between Psophia viridis, P. dextralis and consistentes e distintos para cada um dos táxons, exceto P. obscura, respectively. para P. v. interjecta, cujos caracteres indicados pelos Taxonomic research can also be an important autores como diagnósticos são resultado de variação tool for conservation of the Dark-winged Trum- individual. Não foi observada variação clinal ou sinal peters. While subspecies are not often considered de intergradação entre estes táxons, mesmo nas regiões when addressing the issue of threatened taxa, Pso- próximas às cabeceiras dos grandes rios amazônicos, onde phia obscura is in the Brazilian list of threatened supostamente as populações poderiam entrar em contato. birds (as P. v. obscura, IBAMA, 2003). The Belém Sugere-se que as subespécies correntemente aceitas sejam center of endemism, where P. obscura is found, is an elevadas ao nível de espécie, a saber: Psophia viridis area with many endemic and/or threatened birds, Spix, 1825, que ocorre no interflúvio Madeira-Tapajós; such as the Golden Parakeet (Guarouba guarouba), P. dextralis Conover, 1934, encontrado no interflúvio among others (Roma, 1996; Silveira & Belmonte, Tapajós-Tocantins, e P. obscura Pelzeln, 1857, que 2005) and is the most degraded and logged area distribui-se da margem direita do rio Tocantins até o in the Brazilian Amazon. Trumpeters require quite oeste do Estado do Maranhão. large home ranges and are sensitive to hunting, and hence are much more likely to disappear in Palavras-chave: Psophiidae; Psophia; Psophia viridis; disturbed sites. The main conservation strategy for Taxonomia. 554 Oppenheimer, M. & Silveira, L.F.: Taxonomy of Dark-winged Trumpeter

Acknowledgments Novaes, F.C. & Lima, M.F.C. 1991. As aves do rio Peixoto de Azevedo, Mato Grosso, Brasil. Revista Brasileira de Zoologia, 7(3):351‑381. We thank the Conselho Nacional de Desen- Oppenheimer, M. 2008. Psophia viridis obscura Pelzeln, 1857. In: Machado, A.B.M; Drummond, G.M. & Paglia, A.P. volvimento Científico – CNPq – and the Programa de (Eds.), Livro vermelho da fauna brasileira ameaçada de extinção. Apoio à Pós-Graduação (PROAP) for financial support. Ministério do Meio Ambiente, Brasília; Fundação Biodiversitas, We also appreciate the help and collaboration of the Belo Horizonte. (Série Biodiversidade, 19). curators at the Brazilian ornithological collections for Oren, D.C. 1990. New and Reconfirmed Bird Records from the State of Maranhão, Brazil. Goeldiana Zoologia, (4):1‑13. providing access to the collections, especially Alexandre Oren, D.C. 1991. Aves do Estado do Maranhão, Brasil. Goeldiana Aleixo (MPEG) and Marcos Raposo (MNRJ) and our Zoologia, (9):1‑55. colleagues at MPEG, MNRJ, USP and MZUSP from Paynter Jr., R.A. & Traylor, M.A. 1991. Ornithological gazetteer their support. We also thank the curators of foreign or- of Brazil. Museum of Comparative Zoology, Cambridge, 2v., 788 p. nithological collections for their help with information Pelzeln, A. von. 1857. Psophia obscura. Sitzungsbericht about the type specimens. We would like to give special mathematisch.-naturwissenschaften. Classe Akademie der thanks to Prof. Dra. Elizabeth Höfling, Prof. Dr. Her- Wissenschaften, Wien, 24:373. culano Alvarenga, Prof. Dr. Miguel Trefaut Rodrigues, Pelzeln, A. von. 1871. Zur Ornithologie brasiliens: Resultate von Johann Natterers Reisen in den Jahren 1817 bis 1835. A. and Vitor Q. Piacentini. An anonymous reviewer Pichler’s Witwe & Sohn, Viena, 462 p. makes important suggestions in the manuscript. Peters, J.L. 1934. Check-list of birds of the world. Harvard University Press, Cambridge, v. 2. Pinto, O.M.O. 1938. Catálogo das aves do Brasil e lista dos exemplares que as representam no Museu Paulista (1ª parte). References Revista do Museu Paulista, 22:1‑566. Pinto, O.M.O. 1978. Novo Catálogo das Aves do Brasil. Empresa Baldwin, S.P.; Oberholser, H.C. & Worley, L.G. 1931. Measurements Gráfica da Revista dos Tribunais, São Paulo, v. 1. of birds. Cleveland Museum of Natural History, Cleveland. Roma, J.C. 1996. Composição e vulnerabilidade da avifauna do Blake, E.R. 1977. Manual of Neotropical Birds: Spheniscidae leste do Estado do Pará, Brasil. (Dissertação de Mestrado). (Penguins) to Laridae (Gulls and Allies). University of Chicago Universidade Federal do Pará, Belém. Press, Chicago, v.1, 674 p. Schauensee, R.M. 1970. A guide to the birds of South America. Brabourne, L. & Chubb, C. 1912. The Birds of South America. Oliver & Boyd, Edinburgh. R.H. Porter, London, v.1, 1.504 p. Sharpe, R.B. 1894. Catalogue of the Fulicariae and Alectorides, in Conover, H.B. 1934. A new trumpeter from Brazil. Proceedings of the collection of the British Museum. British Museum (Natural the Biological Society of Washington, 47:119‑120. History), London, v. 13, 354 p. ESRI (Environmental Systems Research Institute). 2002. ArcView 3.3. Sherman, P.T. 1996. Family Psophiidae (trumpeters). In: Del Graves, G.R. & Zusi, R.L. 1986. Preliminary survey of the avifauna of Hoyo, J.; Elliott, A. & Sargatal, J. (Eds), Handbook of the birds the east bank of the lower Rio Xingu. [Relatório não publicado]. of the world. Lynx Edicions, Barcelona, v. 3, p. 96‑107. Griscom, L. & Greenway, J.C. 1937. Psophia viridis interjecta. Bulletin Sick, H. 1997. Ornitologia Brasileira. 3. ed. Nova Fronteira, Rio of the Museum of Comparative Zoology, Harvard, 81(2):419. de Janeiro, 912 p. Gyldenstolpe, N. 1945. The bird fauna of Rio Juruá in Western Silveira, L.F. & Belmonte, F. 2005. Comportamento reprodutivo Brazil. Kungliga Svenska Vetenskaps akademiens Handlingar, e hábitos da ararajuba, Guarouba guarouba, no município de 22(3):1‑338. Tailândia, Pará. Ararajuba, 13(1):89‑93. Haffer, J. 1974. Avian speciation in tropical South America. Smithe, F.B. 1975. Naturalist’s Color Guide. American Museum of Publications of Nuttall Ornithological Club, 14:1‑390. Natural History, New York. Helbig, A.J.; Knox, A.G.; Parkin, D.T.; Sangster, G. & Snethlage, E. 1914. Catálogo das aves amazonicas: contendo Collinson, M. 2002. Guidelines for assigning species rank. todas as especies descriptas e mencionadas até 1913. Boletim Ibis, 144:518‑525. do Museu Paraense Emilio Goeldi, 8(1911‑12):1‑530. Hellmayr, C.E. & Conover, B. 1942. Catalogue of birds of the Spix, J.B. von. 1825. Psophia viridis. In: Avium species novae quas Americas and the adjacent islands in Field Museum of Natural in itinere per Braziliam – annis 1817‑20 collegit et descripsit. History. Publications of Field Museum of Natural History, Impensis, Monachii, v. 2, p. 66. 13(1):308‑314. Stotz, D.F. 1986. Report of field work at Cachoeira Nazaré, Rio Jiparaná, IBAMA (Instituto Brasileiro do Meio Ambiente e dos Rondonia, Brasil, May to July 1986. [Relatório não publicado]. Recursos Naturais Renováveis). 2003. Lista das Espécies Vanzolini, P.E. 1992. A supplement to the Ornithological Gazetteer da Fauna Brasileira Ameaçadas de Extinção. Anexo à Instrução of Brazil. Museu de Zoologia, Universidade de São Paulo, São Normativa Nº 3, de 27 de maio de 2003 do Ministério do Paulo, 252 p. Meio Ambiente. Disponível em: www.ibama.gov.br/fauna. Vanzolini, P.E. 1993. As viagens de Johann Natterer no Brasil, Acesso em: 30/Maio/2005. 1817‑1835. Papéis Avulsos de Zoologia, 38(3):17‑60.

Linnaeus, C. 1758. Psophia. In: Linnaeus, C. Systema naturae per ZUSP

o M

regna tria naturae, secundum classes, ordines, genera, species, cum Recebido em: 18.08.2009 d characteribus, differentiis… 10. ed. L. Salvii, Holmiae, p. 154. Aceito em: 07.10.2009 Munsell. 1994. Soil color charts. Revised edition. Macbeth Division of Impresso em: 18.12.2009 Kollmorgan Instruments Corporation, New Windsor, New York.

Publicações

Naumburg, E.M.B. 1930. Birds of Matto Grosso, Brazil: A report de on the birds secured by the Roosevelt-Rondon expedition.

Bulletin of the American Museum of Natural History, 60:83‑84. Seção Papéis Avulsos de Zoologia, 49(41), 2009 555

Appendix: Specimens Examined (for abbreviations, see text)

Psophia viridis (n = 24): MZUSP 709, ♂, Aripuanã, AM; MZUSP 10938, ♂, Parintins, AM; MZUSP 17764, ♂, Lago do Baptista, AM; MZUSP 21855, ♂, Lago do Baptista, AM; MZUSP 62339, ♀, Rio Aripuanã, Periquito, AM; MZUSP 76728, ♂, Paca, rio Abacaxis, AM; MPEG 13749, ♀, Vila Braga, rio Tapajós, PA; MPEG 39336, ♀, Cachoeira Nazaré, rio Ji-Paraná, RO; MPEG 39337, ♂, Cachoeira Nazaré, rio Ji-Paraná, RO; MPEG MG 58403, ♀, Itaituba, BR 230, km 64, PA; MPEG 58655, ♂, Traíra-Chororó, AM; MNRJ 9637, ind., Jamari, RO; MNRJ 9640, ♂, Vila Braga, rio Tapajós, PA; MNRJ 9644, ind., Mata do Pirocoluina, Águas do Guaporé, RO; MNRJ 9645, ind., Machado, MT; MNRJ 20557, ♂, Lago do Baptista, AM; MNRJ 20559, ♂, Lago do Baptista, AM; MNRJ 20560, ♂, Igarapé do Arary, AM; MNRJ 20561, ♂, Amazonas, AM; MNRJ 20563, ♂, Lago do Baptista, AM; MNRJ 20564, ♂, Igarapé do Arary, AM; MNRJ 20565, ♀, Lago do Baptista, AM; MNRJ 32872, ♀, Jacareacanga, PA; ZSM B11, Parintins, AM (Type).

Psophia dextralis (n = 73): MZUSP 10610, ♂, Taperinha, PA; MZUSP 10611, ♂, Taperinha, PA; MZUSP 20903, ♂, Piquiatuba, PA; MZUSP 20904, ♀, Bom Jardim, PA; MZUSP 20905, ♀, Caxiricatuba, PA; MZUSP 20461, ♂, Bom Jardim, PA; MZUSP 21272, ♀, Piquiatuba, PA; MZUSP 21311, ♂, Piquiatuba, PA; MZUSP 21413, ♂, Piquiatuba, PA; MZUSP 21419, ♀, Caxiricatuba, PA; MZUSP 21762, ♀, Caxiri- catuba, PA; MZUSP 21766, ♂, Caxiricatuba, PA; MZUSP 21800, ♀, Bom Jardim, PA; MZUSP 21801, ♂, Piquiatuba, PA; MZUSP 21820, ♂, Caxiricatuba, PA; MZUSP 21821, ♀, Caxiricatuba, PA; MZUSP 21822, ♂, Piquiatuba, PA; MZUSP 21823, ♀, Caxiricatuba, PA; MZUSP 21858, ♀, Caxiricatuba, PA; MZUSP 21859, ♀, Caxiricatuba, PA; MZUSP 21860, ♂, Caxiricatuba, PA; MZUSP 21923, ♂, Bom Jardim, PA; MZUSP 21950, ♂, Caxiricatuba, PA; MZUSP 21951, ♂, Caxiricatuba, PA; MZUSP 21952, ♂, Piquiatuba, PA; MZUSP 21953, ♂, Piquiatuba, PA; MZUSP 21984, ♀, Caxiricatuba, PA; MZUSP 22001, ♀, Caxirica- tuba, PA; MZUSP 22098, ♂, Caxiricatuba, PA; MZUSP 22345, ♀, foz do Rio Curuá, PA; MZUSP 22502, ♂, Bom Jardim, PA; MZUSP 32918, ♂, Rio Tapajos, PA; MZUSP 46278, ♀, Fordlandia, PA; MZUSP 46279, ♂, Tapaiuna, PA; MZUSP 46280, ♂, Urucurituba, PA; MZUSP 46281, ind., Fordlandia, PA; MZUSP 58106, ind, Fordlandia, PA; MZUSP 58107, ♂, Fordlandia, PA; MZUSP 58108, ♂, Fordlandia, PA; MZUSP 58109, ♀, Tauari, PA; MZUSP 58110, ♀, Tauari, PA; MZUSP 58111, ♀, Fordlandia, PA; MZUSP 58507, ♀, Ford- landia, PA; MZUSP 58508, ♀, Fordlandia, PA; MPEG 14781, ♂, Rio Sete de Setembro, Posto Garapu, MT; MPEG 26398, ♂, Caxiricatuba, PA; MPEG 37204, ♂, Carajás, Serra Norte, PA; MPEG 37205, ♀, Carajás, Serra Norte, Mata da Lixeira, PA; MPEG 37970, ♂, Marabá, Reserva CVRD, Rio Sororó, PA; MPEG MG 40707, ♀, R. Pracupy, Portel, PA; MPEG MG 40708, ♂, Portel, PA; MPEG MG 40709, ♂, Portel, PA; MPEG 48495, ♂, Santana do Araguaia, Faz. Barra das Princesas, PA; MPEG 48496, ♂, Santana do Araguaia, Faz. Far- tura, PA; MPEG 51281, ♂, Alta Floresta, Rio Teles Pires, boca do Rio Cristalino, MT; MPEG 51282, ♀, Rio Cristalino, marg. direita, 15km acima do Teles Pires, MT; MPEG 51283, ♀, Alta Floresta, Rio Cristalino, marg. direita, 15km acima do Rio Teles Pires, MT; MPEG 51284, ♂, Alta Floresta, Rio Teles Pires, marg. esquerda, frente boca Rio Cristalino, MT; MPEG 58592, ind, Serra do Cachimbo, Base Aeronáutica, PA; MNRJ 20566, ♂, Caxiricatuba, PA; MNRJ 25249, ♂, Curuá-tinga, afluente do Curuá-una, PA; MNRJ 25251, ♂, Curua- tinga, afluente do Curua-una, PA; MNRJ 32866, ♂, Jacaré, baixo Culuene (Xingu), MT; MNRJ 32867, ♂, Jacaré, baixo Culuene (Xingu), MT; MNRJ 32868, ♀, Jacaré, baixo Culuene (Alto Xingu), MT; MNRJ 32869, ♀, Jacaré, baixo Culuene (Alto Xingu), MT; MNRJ 32870, ♀, Jacare, baixo Culuene, MT; MNRJ 32871, ♀, Diauarum, Alto Xingu, MT; MNRJ 32873, ♂, Cururú-assu; MNRJ 32874, ♀, Alto Cururu; MNRJ 32875, ♀, Alto Cururu; MNRJ 32876, ♀, Alto Cururu; FMNH 410480, ♂, Tauari, PA (Type).

Psophia obscura (n = 11): MZUSP 43898, ♂, Capim, BR 14, km 93, PA; MZUSP 43899, ♂, Capim, BR 14, km 93, PA; MPEG 1697, ♂, Rio Acará, PA; MPEG 1698, ♂, Rio Acará, PA; MPEG 6584, ♂, Jardim Zoológi- co; MPEG 14440, ♀, Rodovia Belem-Brasília, km 75; MPEG 32003, ♂, Ourem, Sitio Fé em Deus; MPEG 32002, ♂, Mun. Ourem, Sítio Fé em Deus; MPEG 32376, ♂, Ourem, Igarape Pedral, PA; MPEG 36328, ind., Tucuruí, Jacundazinho, PA; MPEG 37338, ind., Buriticupu, MA.